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The power of genetic diversity in genome-wide association studies of lipids

Nature volume 600, pages 675–679 (2021)Cite this article

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Abstract

Increased blood lipid levels are heritable risk factors of cardiovascular disease with varied prevalence worldwide owing to different dietary patterns and medication use1. Despite advances in prevention and treatment, in particular through reducing low-density lipoprotein cholesterol levels2, heart disease remains the leading cause of death worldwide3. Genome-wideassociation studies (GWAS) of blood lipid levels have led to important biological and clinical insights, as well as new drug targets, for cardiovascular disease. However, most previous GWAS4,5,6,7,8,9,10,11,12,13,14,15,16,17,18,19,20,21,22,23 have been conducted in European ancestry populations and may have missed genetic variants that contribute to lipid-level variation in other ancestry groups. These include differences in allele frequencies, effect sizes and linkage-disequilibrium patterns24. Here we conduct a multi-ancestry, genome-wide genetic discovery meta-analysis of lipid levels in approximately 1.65 million individuals, including 350,000 of non-European ancestries. We quantify the gain in studying non-European ancestries and provide evidence to support the expansion of recruitment of additional ancestries, even with relatively small sample sizes. We find that increasing diversity rather than studying additional individuals of European ancestry results in substantial improvements in fine-mapping functional variants and portability of polygenic prediction (evaluated in approximately 295,000 individuals from 7 ancestry groupings). Modest gains in the number of discovered loci and ancestry-specific variants were also achieved. As GWAS expand emphasis beyond the identification of genes and fundamental biology towards the use of genetic variants for preventive and precision medicine25, we anticipate that increased diversity of participants will lead to more accurate and equitable26 application of polygenic scores in clinical practice.

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Fig. 1: Comparison of identified loci across ancestry groups.
Fig. 2: Inclusion of multiple ancestries drives improved fine-mapping.
Fig. 3: Multi-ancestry LDL-C PRS show similar performance across ancestry groups.

Data availability

The GWAS meta-analysis results (including both ancestry-specific and multi-ancestry analyses) and risk score weights are available at http://csg.sph.umich.edu/willer/public/glgc-lipids2021. The optimized multi-ancestry and single-ancestry PRS weights are deposited in the PGS Catalogue (https://www.pgscatalog.org/) accession numbers PGS000886–PGS000897 (all intervening numbers).

Code availability

The code EasyQC is available at www.genepi-regensburg.de/easyqc, and Raremetal is available at https://github.com/SailajaVeda/raremetal.

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Acknowledgements

Funding for the Global Lipids Genetics Consortium was provided by the NIH (R01-HL127564). This research was conducted using the UK Biobank Resource under application number 24460. Computing support and file management for central meta-analysis by S. Caron is acknowledged. This research is based on data from the MVP, Office of Research and Development, Veterans Health Administration, and was supported by awards 2I01BX003362-03A1 and 1I01BX004821-01A1. This publication does not represent the views of the Department of Veteran Affairs or the United States Government. Study-specific acknowledgements are provided in the Supplementary Information.

Author information

Author notes

  1. These authors contributed equally: Shoa L. Clarke, Kuan-Han H. Wu, Stavroula Kanoni, Greg J. M. Zajac, Shweta Ramdas

  2. These authors jointly supervised this work: Gina M. Peloso, Christopher D. Brown, Andrew P. Morris, Themistocles L. Assimes, Panos Deloukas, Yan V. Sun, Cristen J. Willer

Authors and Affiliations

  1. Department of Internal Medicine, Division of Cardiology, University of Michigan, Ann Arbor, MI, USA

    Sarah E. Graham, Ida Surakka, Chao Xue, Jifeng Zhang, Y. Eugene Chen & Cristen J. Willer

  2. VA Palo Alto Health Care System, Palo Alto, CA, USA

    Shoa L. Clarke, Xiang Zhu, Philip S. Tsao & Themistocles L. Assimes

  3. Department of Medicine, Division of Cardiovascular Medicine, Stanford University School of Medicine, Stanford, CA, USA

    Shoa L. Clarke, Erik Ingelsson, Philip S. Tsao & Themistocles L. Assimes

  4. Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, MI, USA

    Kuan-Han H. Wu, Wei Zhou & Cristen J. Willer

  5. William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London, UK

    Stavroula Kanoni, Eirini Marouli, Olga Giannakopoulou, Konain Fatima Bhatti & Panos Deloukas

  6. Department of Biostatistics and Center for Statistics Genetics, University of Michigan, Ann Arbor, MI, USA

    Greg J. M. Zajac, Anne U. Jackson & Michael Boehnke

  7. Department of Genetics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA

    Shweta Ramdas, Anurag Verma & Christopher D. Brown

  8. Clinical Pharmacology, William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, UK

    Ioanna Ntalla

  9. Endocrinology, Boston Children’s Hospital, Boston, MA, USA

    Sailaja Vedantam & Joel N. Hirschhorn

  10. Medical and Population Genetics, Broad Institute, Cambridge, MA, USA

    Sailaja Vedantam & Joel N. Hirschhorn

  11. Department of Genetic Epidemiology, University of Regensburg, Regensburg, Germany

    Thomas W. Winkler, Mathias Gorski, Martina E. Zimmermann, Iris M. Heid & Klaus J. Stark

  12. McDonnell Genome Institute and Department of Medicine, Washington University, St Louis, MO, USA

    Adam E. Locke

  13. Department of Precision Medicine, Division of Genome Science, National Institute of Health, Cheongju-si, South Korea

    Mi Yeong Hwang, Sohee Han, Hye-Mi Jang, Kyungheon Yoon, Young Jin Kim & Bong-Jo Kim

  14. Tohoku Medical Megabank Organization, Tohoku University, Sendai, Japan

    Akira Narita, Gen Tamiya & Masayuki Yamamoto

  15. Sydney Brenner Institute for Molecular Bioscience, Faculty of Health Sciences, University of the Witwatersrand, Johannesburg, South Africa

    Ananyo Choudhury, Scott Hazelhurst & Michèle Ramsay

  16. Center for Research on Genomics and Global Health, National Human Genome Research Institute, National Institutes of Health, Bethesda, MD, USA

    Amy R. Bentley, Kenneth Ekoru & Charles N. Rotimi

  17. Blizard Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London, UK

    Bhavi Trivedi, Karen A. Hunt & David A. van Heel

  18. Wellcome Sanger Institute, Hinxton, UK

    Hilary C. Martin & Nigel W. Rayner

  19. Department of Epidemiology, Emory University Rollins School of Public Health, Atlanta, GA, USA

    Qin Hui & Yan V. Sun

  20. Atlanta VA Health Care System, Decatur, GA, USA

    Qin Hui & Yan V. Sun

  21. Malcolm Randall VA Medical Center, Gainesville, FL, USA

    Derek Klarin

  22. Division of Vascular Surgery and Endovascular Therapy, University of Florida College of Medicine, Gainesville, FL, USA

    Derek Klarin

  23. Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA

    Derek Klarin & Masahiro Kanai

  24. Department of Statistics, The Pennsylvania State University, University Park, PA, USA

    Xiang Zhu

  25. Huck Institutes of the Life Sciences, The Pennsylvania State University, University Park, PA, USA

    Xiang Zhu

  26. Department of Statistics, Stanford University, Stanford, CA, USA

    Xiang Zhu

  27. deCODE genetics/Amgen, Reykjavik, Iceland

    Gudmar Thorleifsson, Anna Helgadottir, Daniel F. Gudbjartsson, Hilma Holm, Unnur Thorsteinsdottir & Kari Stefansson

  28. School of Engineering and Natural Sciences, University of Iceland, Reykjavik, Iceland

    Daniel F. Gudbjartsson

  29. Department of Clinical Biochemistry, Landspitali–National University Hospital of Iceland, Reykjavik, Iceland

    Isleifur Olafsson

  30. Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan

    Masato Akiyama, Saori Sakaue, Masahiro Kanai, Yoichiro Kamatani & Yukinori Okada

  31. Department of Ophthalmology, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan

    Masato Akiyama

  32. Department of Statistical Genetics, Osaka University Graduate School of Medicine, Osaka, Japan

    Saori Sakaue & Yukinori Okada

  33. Department of Allergy and Rheumatology, Graduate School of Medicine, The University of Tokyo, Tokyo, Japan

    Saori Sakaue

  34. Laboratory for Statistical and Translational Genetics, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan

    Chikashi Terao

  35. Department of Biomedical Informatics, Harvard Medical School, Boston, MA, USA

    Masahiro Kanai

  36. Analytic and Translational Genetics Unit, Massachusetts General Hospital, Boston, MA, USA

    Wei Zhou

  37. Program in Medical and Population Genetics, Broad Institute of Harvard and MIT, Cambridge, MA, USA

    Wei Zhou & Pradeep Natarajan

  38. K.G. Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, NTNU, Norwegian University of Science and Technology, Trondheim, Norway

    Ben M. Brumpton, Humaira Rasheed, Kristian Hveem & Bjørn Olav Åsvold

  39. MRC Integrative Epidemiology Unit, University of Bristol, Bristol, UK

    Ben M. Brumpton & Humaira Rasheed

  40. Department of Thoracic Medicine, St Olavs Hospital, Trondheim University Hospital, Trondheim, Norway

    Ben M. Brumpton

  41. Institute for Molecular Medicine Finland (FIMM), HiLIFE, University of Helsinki, Helsinki, Finland

    Sanni E. Ruotsalainen, Aki S. Havulinna, Anu Loukola, Sailalitha Bollepalli, Jaakko Kaprio, Samuli Ripatti & Elisabeth Widén

  42. Department of Public Health and Welfare, Finnish Institute for Health and Welfare, Helsinki, Finland

    Aki S. Havulinna, Pekka Jousilahti & Veikko Salomaa

  43. Department of Genetics, Institute for Biomedical Informatics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA

    Yogasudha Veturi, Yuki Bradford, Jason E. Miller & Marylyn D. Ritchie

  44. Division of Clinical Pharmacology, Department of Medicine, Vanderbilt University Medical Center, Nashville, TN, USA

    QiPing Feng

  45. Department of Medicine (Medical Genetics), University of Washington, Seattle, WA, USA

    Elisabeth A. Rosenthal

  46. Division of Biomedical Informatics, Cincinnati Children’s Hospital Medical Center, Cincinnati, OH, USA

    Todd Lingren

  47. Center for Genetic Medicine, Northwestern University, Evanston, IL, USA

    Jennifer Allen Pacheco

  48. Genentech, South San Francisco, CA, USA

    Sarah A. Pendergrass & Mark I. McCarthy

  49. Fred Hutchinson Cancer Research Center, Division of Public Health Sciences, Seattle, WA, USA

    Jeffrey Haessler & Charles Kooperberg

  50. Division of Preventive Medicine, Brigham and Women’s Hospital, Boston, MA, USA

    Franco Giulianini, Julie E. Buring, Paul M. Ridker & Daniel I. Chasman

  51. Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, UK

    Archie Campbell

  52. Usher Institute for Population Health Sciences and Informatics, The University of Edinburgh, Edinburgh, UK

    Archie Campbell

  53. Clinical Trial Service Unit and Epidemiological Studies Unit, Nuffield Department of Population Health, University of Oxford, Oxford, UK

    Kuang Lin, Iona Y. Millwood, Zhengming Chen, Robin G. Walters & Michael V. Holmes

  54. Medical Research Council Population Health Research Unit, Nuffield Department of Population Health, University of Oxford, Oxford, UK

    Iona Y. Millwood, Zhengming Chen, Robin G. Walters & Michael V. Holmes

  55. Department of Population Medicine, Qatar University College of Medicine, QU Health, Doha, Qatar

    George Hindy

  56. Center for Non-Communicable Diseases, Karachi, Pakistan

    Asif Rasheed, Shahid Abbas & Danish Saleheen

  57. Survey Research Center, Institute for Social Research, University of Michigan, Ann Arbor, MI, USA

    Jessica D. Faul, David R. Weir & Jennifer A. Smith

  58. Department of Epidemiology, School of Public Health, University of Michigan, Ann Arbor, MI, USA

    Wei Zhao

  59. Program in Genetic Epidemiology and Statistical Genetics, Department of Epidemiology, Harvard T. H. Chan School of Public Health, Boston, MA, USA

    Constance Turman, Hongyan Huang & Peter Kraft

  60. Department of Epidemiology, University of North Carolina, Chapel Hill, NC, USA

    Mariaelisa Graff, Heather M. Highland, Kristin L. Young & Kari E. North

  61. Wellcome Centre for Human Genetics, University of Oxford, Oxford, UK

    Anubha Mahajan, Anuj Goel, Nigel W. Rayner, Mark I. McCarthy & Hugh Watkins

  62. Human Genetics Center, Department of Epidemiology, Human Genetics, and Environmental Sciences, School of Public Health, The University of Texas Health Science Center at Houston, Houston, TX, USA

    Michael R. Brown, Paul S. de Vries & Alanna C. Morrison

  63. Department of Epidemiology and Biostatistics, Imperial College London, London, UK

    Weihua Zhang & John C. Chambers

  64. Department of Cardiology, Ealing Hospital, London North West University Healthcare NHS Trust, Southall, UK

    Weihua Zhang, John C. Chambers & Jaspal S. Kooner

  65. Imperial College Healthcare NHS Trust, London, UK

    Weihua Zhang & Jaspal S. Kooner

  66. Department of Biostatistics, Center for Statistical Genetics, University of Michigan, Ann Arbor, MI, USA

    Ketian Yu, Ellen M. Schmidt, Anita Pandit, Xianyong Yin & Goncalo Abecasis

  67. Department of Medical Sciences, Molecular Epidemiology and Science for Life Laboratory, Uppsala University, Uppsala, Sweden

    Stefan Gustafsson & Erik Ingelsson

  68. MRC Epidemiology Unit, University of Cambridge School of Clinical Medicine, Cambridge, UK

    Jian’an Luan, Nicholas J. Wareham & Claudia Langenberg

  69. Cardiovascular Epidemiology Unit, Department of Public Health and Primary Care, University of Cambridge Strangeways Research Laboratory, Cambridge, UK

    Jing-Hua Zhao

  70. Center for Genomic Medicine, Kyoto University Graduate School of Medicine, Kyoto, Japan

    Fumihiko Matsuda, Takahisa Kawaguchi & Yasuharu Tabara

  71. Department of Internal Medicine, Erasmus MC, University Medical Center Rotterdam, Rotterdam, The Netherlands

    Carolina Medina-Gomez & Joyce B. J. van Meurs

  72. Department of Epidemiology, Erasmus MC, University Medical Center Rotterdam, Rotterdam, The Netherlands

    Carolina Medina-Gomez, Natalie Terzikhan, Hieab H. H. Adams, M. Arfan Ikram, Joyce B. J. van Meurs & Mohsen Ghanbari

  73. Department of Biostatistics, Boston University School of Public Health, Boston, MA, USA

    Achilleas Pitsillides, L. Adrienne Cupples & Gina M. Peloso

  74. Department of Biological Psychology, Behavioral and Movement Sciences, Vrije Universiteit Amsterdam, Amsterdam, The Netherlands

    Jouke Jan Hottenga, Gonneke Willemsen, Dorret I. Boomsma & Eco J. C. de Geus

  75. Amsterdam Public Health, VU Medical Center Amsterdam, Amsterdam, The Netherlands

    Jouke Jan Hottenga

  76. Amsterdam Public Health Research Institute, VU Medical Center Amsterdam, Amsterdam, The Netherlands

    Gonneke Willemsen

  77. Genetics of Complex Traits, University of Exeter Medical School, University of Exeter, Exeter, UK

    Andrew R. Wood, Yingji Ji & Timothy M. Frayling

  78. Department of Clinical Acupuncture and Moxibustion, Nanjing University of Chinese Medicine, Nanjing, China

    Zishan Gao

  79. Research Unit of Molecular Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany

    Zishan Gao & Christian Gieger

  80. Institute of Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany

    Zishan Gao, Annette Peters & Christian Gieger

  81. MRC Integrative Epidemiology Unit at the University of Bristol, Bristol, UK

    Simon Haworth, Ruth E. Mitchell & Nicholas J. Timpson

  82. Bristol Dental School, University of Bristol, Bristol, UK

    Simon Haworth

  83. Population Health Sciences, Bristol Medical School, University of Bristol, Bristol, UK

    Ruth E. Mitchell & Nicholas J. Timpson

  84. Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore, Singapore

    Jin Fang Chai, Xueling Sim, E. Shyong Tai & Rob M. van Dam

  85. Center for Clinical Research and Prevention, Bispebjerg and Frederiksberg Hospital, Copenhagen, Denmark

    Mette Aadahl, Line L. Kårhus, Line T. Møllehave, Betina H. Thuesen, Allan Linneberg & Thomas M. Dantoft

  86. The Institute for Translational Genomics and Population Sciences, Department of Pediatrics, Lundquist Institute for Biomedical Innovations (Formerly LABioMed) at Harbor-UCLA Medical Center, Torrance, CA, USA

    Jie Yao, Xiaohui Li, Yii-Der Ida Chen, Xiuqing Guo & Jerome I. Rotter

  87. Center for Public Health Genomics, University of Virginia, Charlottesville, VA, USA

    Ani Manichaikul

  88. William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, John Vane Science Centre, Queen Mary University of London, London, UK

    Helen R. Warren, Julia Ramirez, Mark J. Caulfield & Patricia B. Munroe

  89. NIHR Barts Cardiovascular Biomedical Research Centre, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, UK

    Helen R. Warren, Morris Brown, Mark J. Caulfield, Patricia B. Munroe & Panos Deloukas

  90. Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

    Jette Bork-Jensen, Henrik Vestergaard, Oluf Pedersen, Torben Hansen & Niels Grarup

  91. Division of Cardiovascular Medicine, Radcliffe Department of Medicine, John Radcliffe Hospital, University of Oxford, Oxford, UK

    Anuj Goel

  92. Group of Genomics of Complex Diseases, Research Institute of Hospital de la Santa Creu i Sant Pau (IIB Sant Pau), Barcelona, Spain

    Maria Sabater-Lleal

  93. Cardiovascular Medicine Unit, Department of Medicine, Karolinska Institutet, Center for Molecular Medicine, Karolinska University Hospital, Stockholm, Sweden

    Maria Sabater-Lleal

  94. Department of Internal Medicine, Section Gerontology and Geriatrics, Leiden University Medical Center, Leiden, The Netherlands

    Raymond Noordam

  95. Institute for Genetic and Biomedical Research, Italian National Council of Research (IRGB CNR), Cagliari,, Italy

    Carlo Sidore

  96. Institute for Genetic and Biomedical Research, Italian National Council of Research (IRGB CNR), Lanusei, Italy

    Edoardo Fiorillo

  97. University Center for Primary Care and Public Health, University of Lausanne, Lausanne, Switzerland

    Aaron F. McDaid

  98. Swiss Institute of Bioinformatics, Lausanne, Switzerland

    Aaron F. McDaid & Zoltan Kutalik

  99. Department of Medicine, Internal Medicine, Lausanne University Hospital and University of Lausanne, Lausanne, Switzerland

    Pedro Marques-Vidal

  100. Department of Epidemiology and Biostatistics, MRC–PHE Centre for Environment and Health, School of Public Health, Imperial College London, London, UK

    Matthias Wielscher & Marjo-Riitta Jarvelin

  101. Department of Cardiology, Leiden University Medical Center, Leiden, The Netherlands

    Stella Trompet & J. Wouter Jukema

  102. Department of Internal Medicine, Section of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, The Netherlands

    Stella Trompet

  103. BHF Glasgow Cardiovascular Research Centre, Faculty of Medicine, Glasgow, UK

    Naveed Sattar

  104. Institute for Cardiogenetics, University of Lübeck, DZHK (German Research Centre for Cardiovascular Research), Partner site Hamburg/Lübeck/Kiel, University Heart Center Lübeck, Lübeck, Germany

    Matthias Munz

  105. Charité–University Medicine Berlin, Member of Freie Universität Berlin and Humboldt-Universität zu Berlin, Berlin, Germany

    Matthias Munz

  106. Department of Periodontology and Synoptic Dentistry, Berlin Institute of Health, Institute for Dental and Craniofacial Sciences, Berlin, Germany

    Matthias Munz

  107. Deutsches Herzzentrum München, Klinik für Herz- und Kreislauferkrankungen, Technische Universität München, Munich, Germany

    Lingyao Zeng

  108. Deutsches Zentrum für Herz-Kreislauf-Forschung (DZHK), Partner site Munich Heart Alliance, Munich, Germany

    Lingyao Zeng

  109. Key Laboratory of Cardiovascular Epidemiology and Department of Epidemiology, State Key Laboratory of Cardiovascular Disease, Fuwai Hospital, National Center for Cardiovascular Diseases, Chinese Academy of Medical Sciences and Peking Union Medical College, Beijing, China

    Jianfeng Huang, Bin Yang, Shufeng Chen, Fangchao Liu, Xiangfeng Lu & Dongfeng Gu

  110. Lund University Diabetes Centre, Malmo, Sweden

    Alaitz Poveda, Azra Kurbasic & Paul W. Franks

  111. Department of Genetics and Pharmacology, Institute of Genetic Epidemiology, Medical University of Innsbruck, Innsbruck, Austria

    Claudia Lamina & Lukas Forer

  112. German Chronic Kidney Disease Study, Innsbruck, Austria

    Claudia Lamina, Lukas Forer & Florian Kronenberg

  113. Institute for Medical Informatics, Statistics and Epidemiology, University of Leipzig, Leipzig, Germany

    Markus Scholz & Markus Loeffler

  114. LIFE Research Centre for Civilization Diseases, University of Leipzig, Leipzig, Germany

    Markus Scholz, Joachim Thiery & Markus Loeffler

  115. Radboud University Medical Center, Radboud Institute for Health Sciences, Nijmegen, The Netherlands

    Tessel E. Galesloot, Lambertus A. L. M. Kiemeney & Jacqueline de Graaf

  116. Quantinuum Research, Wayne, PA, USA

    Jonathan P. Bradfield

  117. Division of Statistical Genomics, Department of Genetics, Washington University School of Medicine, St Louis, MO, USA

    E. Warwick Daw, Mary F. Feitosa, Mary K. Wojczynski, Karen Schwander & Michael A. Province

  118. Department of Epidemiology, University of Pittsburgh, Pittsburgh, PA, USA

    Joseph M. Zmuda

  119. Institute for Biomedicine, Eurac Research, Affiliated Institute of the University of Lübeck, Bolzano, Italy

    Jonathan S. Mitchell, Christian Fuchsberger & Peter P. Pramstaller

  120. Department of Clinical Biochemistry, Lillebaelt Hospital, Vejle, Denmark

    Henry Christensen & Ivan Brandslund

  121. Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, USA

    Jennifer A. Brody & Joshua C. Bis

  122. The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, NY, USA

    Michael Preuss, Girish N. Nadkarni & Ruth J. F. Loos

  123. Department of Twin Research and Genetic Epidemiology, King’s College London, London, UK

    Massimo Mangino, Paraskevi Christofidou & Tim D. Spector

  124. NIHR Biomedical Research Centre at Guy’s and St Thomas’ Foundation Trust, London, UK

    Massimo Mangino

  125. Department of Cardiology, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands

    Niek Verweij, Jan W. Benjamins & Pim van der Harst

  126. Institute of Cardiovascular Sciences, University College London, London, UK

    Jorgen Engmann

  127. Department of Epidemiology and Public Health, University College London, London, UK

    Jorgen Engmann & Mika Kivimaki

  128. Department of Psychiatry, University of Pennsylvania, Philadelphia, PA, USA

    Rachel L. Kember

  129. Amsterdam UMC, Department of Epidemiology and Biostatistics, Amsterdam Public Health Research Institute, Amsterdam, The Netherlands

    Roderick C. Slieker, Morgan O. Obura, Joline W. J. Beulens & Leen M. ’t Hart

  130. Department of Cell and Chemical Biology, Leiden University Medical Center, Leiden, The Netherlands

    Roderick C. Slieker & Leen M. ’t Hart

  131. Montreal Heart Institute, Montreal, Quebec, Canada

    Ken Sin Lo, Jean-Claude Tardif & Guillaume Lettre

  132. Icelandic Heart Association, Reykjavik, Iceland

    Nuno R. Zilhao & Vilmundur Gudnason

  133. Department of Anthropology, University of Toronto at Mississauga, Mississauga, Ontario, Canada

    Phuong Le & Esteban J. Parra

  134. Vth Department of Medicine, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany

    Marcus E. Kleber, Graciela E. Delgado & Winfried März

  135. Synlab MVZ Humangenetik Mannheim, Mannheim, Germany

    Marcus E. Kleber

  136. Biomedical Technology Research Center, Tokushima Research Institute, Otsuka Pharmaceutical, Tokushima, Japan

    Daisuke D. Ikeda, Hiroyuki Iha & Haretsugu Hishigaki

  137. Institute for Molecular Bioscience, The University of Queensland, Brisbane, Queensland, Australia

    Jian Yang

  138. Institute for Advanced Research, Wenzhou Medical University, Wenzhou, China

    Jian Yang

  139. Nuffield Department of Population Health, University of Oxford, Oxford, UK

    Jun Liu & Cornelia M. van Duijn

  140. Laboratory of Neurogenetics, National Institute on Aging, NIH, Bethesda, MD, USA

    Hampton L. Leonard & Mike A. Nalls

  141. Data Tecnica International, Glen Echo, MD, USA

    Hampton L. Leonard & Mike A. Nalls

  142. MRC Human Genetics Unit, Institute of Genetics and Cancer, University of Edinburgh, Western General Hospital, Edinburgh, UK

    Jonathan Marten, Paul R. H. J. Timmers, James F. Wilson, Veronique Vitart & Caroline Hayward

  143. Institute for Medical Informatics, Biometrie and Epidemiology, University of Duisburg-Essen, Essen, Germany

    Börge Schmidt, Marina Arendt & Karl-Heinz Jöckel

  144. Department of Computer Science, University of Applied Sciences and Arts Dortmund, Dortmund, Germany

    Marina Arendt

  145. Centre for Public Health, Queen’s University of Belfast, Belfast, UK

    Laura J. Smyth, Marisa Cañadas-Garre, Amy Jayne McKnight & Frank Kee

  146. Department of Epidemiology and Biostatistics, School of Public Health, Tongji Medical College, Huazhong University of Science and Technology, Wuhan, China

    Chaolong Wang

  147. Genome Institute of Singapore, Agency for Science, Technology and Research, Singapore, Singapore

    Chaolong Wang, Rajkumar Dorajoo, Jianjun Liu & Chiea Chuen Khor

  148. Public Health Informatics Unit, Department of Integrated Health Sciences, Nagoya University Graduate School of Medicine, Nagoya, Japan

    Masahiro Nakatochi

  149. MRC Unit for Lifelong Health and Ageing at UCL, London, UK

    Andrew Wong & Nish Chaturvedi

  150. Department of Pediatrics, Tampere University Hospital, Tampere, Finland

    Nina Hutri-Kähönen

  151. Department of Pediatrics, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland

    Nina Hutri-Kähönen

  152. Brown Foundation Institute of Molecular Medicine, McGovern Medical School, University of Texas Health Science Center at Houston, Houston, TX, USA

    Rui Xia & Myriam Fornage

  153. Departamento de Medicina Genómica y Toxicología Ambiental, Instituto de Investigaciones Biomédicas, UNAM, Ciudad de Mexico, Mexico, Mexico

    Alicia Huerta-Chagoya

  154. Departamento de Genómica Computacional, Instituto Nacional de Medicina Genómica, Ciudad de Mexico, Mexico, Mexico

    Juan Carlos Fernandez-Lopez

  155. Center for Diabetes Research, University of Bergen, Bergen, Norway

    Valeriya Lyssenko

  156. Lund University Diabetes Center, Lund University, Malmo, Sweden

    Valeriya Lyssenko

  157. Genomic Research on Complex Diseases (GRC Group), CSIR–Centre for Cellular and Molecular Biology, Hyderabad, India

    Meraj Ahmed & Giriraj R. Chandak

  158. Epidemiology, School of Public Health, University of Alabama at Birmingham, Birmingham, AL, USA

    Marguerite R. Irvin

  159. Hunter Medical Research Institute, Newcastle, New South Wales, Australia

    Christopher Oldmeadow, John Attia & Rodney J. Scott

  160. Medical Research Institute, Kangbuk Samsung Hospital, Sungkyunkwan University School of Medicine, Seoul, Korea

    Han-Na Kim

  161. Department of Clinical Research Design and Evaluation, SAIHST, Sungkyunkwan University, Seoul, Korea

    Han-Na Kim

  162. Center for Cohort Studies, Total Healthcare Center, Kangbuk Samsung Hospital, Sungkyunkwan University School of Medicine, Seoul, Korea

    Seungho Ryu

  163. Department of Occupational and Environmental Medicine, Kangbuk Samsung Hospital, Sungkyunkwan University School of Medicine, Seoul, Korea

    Seungho Ryu

  164. Centre for Global Health Research, Usher Institute, University of Edinburgh, Edinburgh, UK

    Paul R. H. J. Timmers, Katherine A. Kentistou, Harry Campbell, Peter K. Joshi & James F. Wilson

  165. Thurston Arthritis Research Center, University of North Carolina, Chapel Hill, NC, USA

    Liubov Arbeeva, Amanda E. Nelson & Yvonne M. Golightly

  166. Division of Biomedical Informatics and Personalized Medicine, Department of Medicine, Anschutz Medical Campus, University of Colorado, Denver, Aurora, CO, USA

    Leslie A. Lange

  167. Ocular Epidemiology, Singapore Eye Research Institute, Singapore National Eye Centre, Singapore, Singapore

    Xiaoran Chai, Ching-Yu Cheng, Tien-Yin Wong & Charumathi Sabanayagam

  168. Department of Ophthalmology, National University of Singapore and National University Health System, Singapore, Singapore

    Xiaoran Chai

  169. Genomics and Molecular Medicine Unit, CSIR–Institute of Genomics and Integrative Biology, New Delhi, India

    Gauri Prasad

  170. Academy of Scientific and Innovative Research, CSIR–Institute of Genomics and Integrative Biology Campus, New Delhi, India

    Gauri Prasad & Dwaipayan Bharadwaj

  171. Departments of Ophthalmology and Human Genetics, Radboud University Nijmegen Medical Center, Nijmegen, The Netherlands

    Laura Lorés-Motta, Marc Pauper & Anneke I. den Hollander

  172. Vanderbilt Epidemiology Center, Division of Epidemiology, Vanderbilt University Medical Center, Nashville, TN, USA

    Jirong Long, Qiuyin Cai, Xiao-Ou Shu & Wei Zheng

  173. Department of Pediatrics, University of California, San Francisco, Oakland, CA,, USA

    Elizabeth Theusch

  174. National Center for Global Health and Medicine, Tokyo, Japan

    Fumihiko Takeuchi & Norihiro Kato

  175. Department of Genetics, University of North Carolina, Chapel Hill, NC, USA

    Cassandra N. Spracklen & Karen L. Mohlke

  176. Department of Biostatistics and Epidemiology, University of Massachusetts-Amherst, Amherst, MA,, USA

    Cassandra N. Spracklen

  177. Department of Cardiovascular Sciences, University of Leicester, Leicester, UK

    Sophie C. Warner, Christopher P. Nelson, Peter S. Braund & Nilesh J. Samani

  178. NIHR Leicester Biomedical Research Centre, Glenfield Hospital, Leicester, UK

    Sophie C. Warner, Christopher P. Nelson, Peter S. Braund & Nilesh J. Samani

  179. Beijing Institute of Ophthalmology, Beijing Key Laboratory of Ophthalmology and Visual Sciences, Beijing, China

    Ya Xing Wang

  180. Beijing Tongren Eye Center, Beijing Tongren Hospital, Capital Medical University, Beijing, China

    Ya Xing Wang & Wen B. Wei

  181. Institute of Genetics and Biophysics “Adriano Buzzati-Traverso”–CNR, Naples, Italy

    Teresa Nutile, Daniela Ruggiero & Marina Ciullo

  182. IRCCS Neuromed, Pozzilli, Isernia, Italy

    Daniela Ruggiero & Marina Ciullo

  183. Department of Psychiatry, Washington University, St Louis, MO, USA

    Yun Ju Sung

  184. Division of Endocrinology and Metabolism, Tri-Service General Hospital Songshan Branch, Taipei, Taiwan

    Yi-Jen Hung

  185. Rush Alzheimer’s Disease Center, Rush University Medical Center, Chicago, IL, USA

    Jingyun Yang & David A. Bennett

  186. Department of Neurological Sciences, Rush University Medical Center, Chicago, IL, USA

    Jingyun Yang & David A. Bennett

  187. Department of Nephrology, University Hospital Regensburg, Regensburg, Germany

    Mathias Gorski, Bettina Jung & Carsten A. Böger

  188. Department of Medicine, Surgery and Health Sciences, University of Trieste, Trieste, Italy

    Marco Brumat & Giorgia Girotto

  189. Department of Molecular Epidemiology, German Institute of Human Nutrition Potsdam-Rehbruecke, Nuthetal, Germany

    Karina Meidtner, Susanne Jäger & Matthias B. Schulze

  190. German Center for Diabetes Research (DZD), Neuherberg, Germany

    Karina Meidtner, Susanne Jäger, Matthias B. Schulze, Annette Peters & Christian Gieger

  191. Department of Epidemiology, School of Public Health, University of Michigan, Ann Arbor, MI, USA

    Lawrence F. Bielak, Jennifer A. Smith & Sharon L. R. Kardia

  192. Department of Genetics and Bioinformatics, Dasman Diabetes Institute, Kuwait, Kuwait

    Prashantha Hebbar, Fahd Al-Mulla & Thangavel Alphonse Thanaraj

  193. Department of Nutrition and Dietetics, School of Health Science and Education, Harokopio University of Athens, Athens, Greece

    Aliki-Eleni Farmaki & George Dedoussis

  194. Department of Population Science and Experimental Medicine, University College London, London, UK

    Aliki-Eleni Farmaki

  195. Department of Neurology, Clinical Division of Neurogeriatrics, Medical University of Graz, Graz, Austria

    Edith Hofer & Reinhold Schmidt

  196. Institute for Medical Informatics, Statistics and Documentation, Medical University of Graz, Graz, Austria

    Edith Hofer

  197. Department of Bioinformatics and Genomics, University of North Carolina at Charlotte, Charlotte, NC, USA

    Maoxuan Lin

  198. Institute for Maternal and Child Health, IRCCS Burlo Garofolo, Trieste, Italy

    Maria Pina Concas

  199. Institute of Genetic and Biomedical Research, National Research Council of Italy, UOS of Sassari, Sassari, Italy

    Simona Vaccargiu

  200. Department of Epidemiology, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands

    Peter J. van der Most & Harold Snieder

  201. Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland

    Niina Pitkänen, Olli T. Raitakari & Katja Pahkala

  202. Centre for Population Health Research, University of Turku and Turku University Hospital, Turku, Finland

    Niina Pitkänen, Olli T. Raitakari & Katja Pahkala

  203. Sleep Medicine and Circadian Disorders, Brigham and Women’s Hospital, Boston, MA, USA

    Brian E. Cade, Jiwon Lee & Susan Redline

  204. Division of Sleep Medicine, Harvard Medical School, Boston, MA, USA

    Brian E. Cade & Susan Redline

  205. Central Diagnostics Laboratory, Division Laboratories, Pharmacy, and Biomedical Genetics, University Medical Center Utrecht, Utrecht University, Utrecht, The Netherlands

    Sander W. van der Laan, Gerard Pasterkamp & Imo E. Hoefer

  206. Laboratory of Epidemiology and Population Science, National Institute on Aging Intramural Research Program, NIH Biomedical Research Center, NIA, Baltimore, MD, USA

    Kumaraswamy Naidu Chitrala

  207. Department of Functional Genomics, Interfaculty Institute for Genetics and Functional Genomics, University of Greifswald and University Medicine Greifswald, Greifswald, Germany

    Stefan Weiss & Georg Homuth

  208. Oneomics, Bucheon, Korea

    Jong Young Lee

  209. Department of Biomedical Science, Hallym University, Chuncheon, Korea

    Hyeok Sun Choi, Jae Hun Shin & Yoon Shin Cho

  210. Centre for Bone and Arthritis Research, Department of Internal Medicine and Clinical Nutrition, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden

    Maria Nethander, Claes Ohlsson & Dan Mellström

  211. Bioinformatics Core Facility, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden

    Maria Nethander

  212. Institute of Medical Informatics and Statistics, Kiel University, Kiel, Germany

    Sandra Freitag-Wolf

  213. Institute of Translational Genomics, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany

    Lorraine Southam, Nigel W. Rayner & Eleftheria Zeggini

  214. Wellcome Trust Sanger Institute, Hinxton, UK

    Lorraine Southam & Eleftheria Zeggini

  215. Oxford Centre for Diabetes Endocrinology and Metabolism, Oxford, UK

    Nigel W. Rayner

  216. School of Medicine and Public Health, College of Health, Medicine and Wellbeing, University of Newcastle, Newcastle, New South Wales, Australia

    Carol A. Wang & Craig E. Pennell

  217. Center for Geriatrics and Gerontology, Division of Endocrinology and Metabolism, Department of Internal Medicine, Taichung Veterans General Hospital, Taichung, Taiwan

    Shih-Yi Lin

  218. School of Medicine, National Yang-Ming University, Taipei, Taiwan

    Shih-Yi Lin

  219. School of Medicine, National Defense Medical Center, Taipei, Taiwan

    Shih-Yi Lin

  220. Division of Endocrinology and Metabolism, Department of Internal Medicine, Taichung Veterans General Hospital, Taichung, Taiwan

    Jun-Sing Wang

  221. Department of Medicine, School of Medicine, National Yang-Ming University, Taipei, Taiwan

    Jun-Sing Wang

  222. Department of Kinesiology, Université Laval, Quebec, Quebec, Canada

    Christian Couture & Louis Pérusse

  223. Department of Clinical Chemistry, Fimlab Laboratories, Tampere, Finland

    Leo-Pekka Lyytikäinen & Terho Lehtimäki

  224. Department of Clinical Chemistry, Finnish Cardiovascular Research Center–Tampere, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland

    Leo-Pekka Lyytikäinen & Terho Lehtimäki

  225. Department of Cardiology, Heart Center, Tampere University Hospital, Tampere, Finland

    Kjell Nikus

  226. Department of Cardiology, Finnish Cardiovascular Research Center–Tampere, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland

    Kjell Nikus

  227. University of Queensland Diamantina Institute, Translational Research Institute, Brisbane, Queensland, Australia

    Gabriel Cuellar-Partida

  228. Department of Medicine, Bornholms Hospital, Ronne, Denmark

    Henrik Vestergaard

  229. School of Public Health, University of Alabama at Birmingham, Birmingham, AL, USA

    Bertha Hildalgo

  230. Cardiology, Division Heart and Lungs, University Medical Center Utrecht, Utrecht University, Utrecht, The Netherlands

    Jessica van Setten, Folkert W. Asselbergs & Adriaan O. Kraaijeveld

  231. Department of Population and Quantitative Health Sciences, Case Western Reserve University, Cleveland, OH, USA

    Xiaoyin Li & Xiaofeng Zhu

  232. Division of Endocrinology, Ohio State University, Columbus, OH, USA

    Rebecca D. Jackson

  233. Department of Epidemiology, University of Washington, Seattle, WA, USA

    Alexander P. Reiner

  234. School of Medicine and Health Sciences, George Washington University, Washington, DC, USA

    Lisa Warsinger Martin

  235. Department of Epidemiology, School of Public Health, Peking University Health Science Center, Beijing, China

    Liming Li

  236. Institute for Laboratory Medicine, University Hospital Leipzig, Leipzig, Germany

    Joachim Thiery

  237. Laboratory of Epidemiology and Population Sciences, National Institute on Aging, NIH, Baltimore, MD, USA

    Lenore J. Launer

  238. Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, Finland

    Olli T. Raitakari

  239. Department of Environmental and Preventive Medicine, Jichi Medical University School of Medicine, Shimotsuke, Japan

    Sahoko Ichihara

  240. Centre for Population Health Sciences, Usher Institute, University of Edinburgh, Edinburgh, UK

    Sarah H. Wild

  241. Department of Functional Pathology, Shimane University School of Medicine, Izumo, Japan

    Toru Nabika

  242. Department of Pediatrics and Adolescent Medicine, Turku University Hospital and University of Turku, Turku, Finland

    Harri Niinikoski

  243. Department of Physiology, University of Turku, Turku, Finland

    Harri Niinikoski

  244. Faculty of Medicine, University of Split, Split, Croatia

    Ivana Kolcic & Ozren Polasek

  245. Medical Department III–Endocrinology, Nephrology, Rheumatology, University of Leipzig Medical Center, Leipzig, Germany

    Peter Kovacs & Anke Tönjes

  246. Department of Nutrition–Dietetics, Harokopio University, Eleftheriou Venizelou, Athens, Greece

    Tota Giardoglou

  247. Department of Clinical Gene Therapy, Osaka University Graduate School of Medicine, Suita, Japan

    Tomohiro Katsuya

  248. Department of Geriatric and General Medicine, Osaka University Graduate School of Medicine, Suita, Japan

    Tomohiro Katsuya

  249. Department of Vascular Surgery, Division of Surgical Specialties, University Medical Center Utrecht, Utrecht University, Utrecht, The Netherlands

    Dominique de Kleijn & Gert J. de Borst

  250. Corneal Dystrophy Research Institute, Department of Ophthalmology, Yonsei University College of Medicine, Seoul, Korea

    Eung Kweon Kim

  251. Department of Radiology and Nuclear Medicine, Erasmus MC, University Medical Center Rotterdam, Rotterdam, The Netherlands

    Hieab H. H. Adams

  252. Julius Centre for Health Sciences and Primary Care, University Medical Centre Utrecht, Utrecht, The Netherlands

    Joline W. J. Beulens

  253. Second Department of Cardiology, Medical School, National and Kapodistrian University of Athens, Attikon University Hospital, Athens, Greece

    Loukianos S. Rallidis

  254. Center for Vision Research, Department of Ophthalmology and The Westmead Institute, University of Sydney, Sydney, New South Wales, Australia

    Paul Mitchell

  255. Menzies Institute for Medical Research, School of Medicine, University of Tasmania, Hobart, Tasmania, Australia

    Alex W. Hewitt

  256. Centre for Eye Research Australia, University of Melbourne, Melbourne, Victoria, Australia

    Alex W. Hewitt

  257. Department of Clinical Physiology, Tampere University Hospital, Tampere, Finland

    Mika Kähönen

  258. Department of Clinical Physiology, Finnish Cardiovascular Research Center–Tampere, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland

    Mika Kähönen

  259. Institute of Nutrition and Functional Foods (INAF), Université Laval, Quebec, Quebec, Canada

    Louis Pérusse

  260. Pennington Biomedical Research Center, Baton Rouge, LA, USA

    Claude Bouchard

  261. Medical School, Faculty of Health and Medical Sciences, University of Western Australia, Perth, Western Australia, Australia

    Trevor A. Mori

  262. Institute of Epidemiology, Kiel University, Kiel, Germany

    Wolfgang Lieb

  263. Institute of Clinical Molecular Biology, Kiel University, Kiel, Germany

    Andre Franke

  264. Department of Drug Treatment, Sahlgrenska University Hospital, Gothenburg, Sweden

    Claes Ohlsson

  265. Geriatric Medicine, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden

    Dan Mellström

  266. Department of Internal Medicine, Ewha Womans University School of Medicine, Seoul, Korea

    Hyejin Lee

  267. Division of Cancer Control and Population Sciences, UPMC Hillman Cancer Center, University of Pittsburgh, Pittsburgh, PA, USA

    Jian-Min Yuan

  268. Department of Epidemiology, Graduate School of Public Health, University of Pittsburgh, Pittsburgh, PA, USA

    Jian-Min Yuan

  269. Healthy Longevity Translational Research Programme, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore

    Woon-Puay Koh

  270. Singapore Institute for Clinical Sciences, Agency for Science Technology and Research (A*STAR), Singapore, Singapore

    Woon-Puay Koh

  271. Department of Endocrinology and Metabolism, Kyung Hee University School of Medicine, Seoul, Korea

    Sang Youl Rhee & Jeong-Taek Woo

  272. Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany

    Henry Völzke

  273. Laboratory of Epidemiology and Population Science, National Institute on Aging Intramural Research Program, NIH Biomedical Research Center, Baltimore, MD, USA

    Michele K. Evans & Alan B. Zonderman

  274. Paavo Nurmi Centre, Sports and Exercise Medicine Unit, Department of Physical Activity and Health, University of Turku, Turku, Finland

    Katja Pahkala

  275. Interdisciplinary Center Psychopathology and Emotion Regulation (ICPE), University of Groningen, University Medical Center Groningen, Groningen, The Netherlands

    Albertine J. Oldehinkel

  276. Institute of Molecular Genetics, National Research Council of Italy, Pavia, Italy

    Ginevra Biino

  277. Gottfried Schatz Research Center for Cell Signaling, Metabolism and Aging, Medical University of Graz, Graz, Austria

    Helena Schmidt

  278. Local Health Unit Toscana Centro, Firenze, Italy

    Stefania Bandinelli

  279. Institute of Nutritional Science, University of Potsdam, Nuthetal, Germany

    Matthias B. Schulze

  280. Institute for Maternal and Child Health IRCCS “Burlo Garofolo”, Trieste, Italy

    Giorgia Girotto

  281. Department of Nephrology, Diabetology, Rheumatology, Traunstein Hospital, Traunstein, Germany

    Carsten A. Böger

  282. KfH Kidney Center Traunstein, Traunstein, Germany

    Carsten A. Böger

  283. Center for Translational and Systems Neuroimmunology, Department of Neurology, Columbia University Medical Center, New York, NY, USA

    Philip L. De Jager

  284. Program in Medical and Population Genetics, Broad Institute, Cambridge, MA, USA

    Philip L. De Jager

  285. Medical School, National and Kapodistrian University Athens, Athens, Greece

    Vasiliki Mamakou

  286. Dromokaiteio Psychiatric Hospital, Athens, Greece

    Vasiliki Mamakou

  287. Clinical Pharmacology, William Harvey Research Institute, Queen Mary University of London, London, UK

    Morris Brown

  288. Department of Ophthalmology, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany

    Jost B. Jonas

  289. Beijing Institute of Ophthalmology, Beijing Key Laboratory of Ophthalmology and Visual Sciences, Beijing Tongren Eye Center, Beijing Tongren Hospital, Capital Medical University, Beijing, China

    Jost B. Jonas

  290. Institute of Molecular and Clinical Ophthalmology Basel, Basel, Switzerland

    Jost B. Jonas

  291. Harvard Medical School, Boston, MA, USA

    Daniel I. Chasman

  292. Harvard Medical School, Boston, MA, USA

    Josep M. Mercader, Julie E. Buring & Paul M. Ridker

  293. Department of Human Genetics, David Geffen School of Medicine at UCLA, University of California, Los Angeles, Los Angeles, CA, USA

    Päivi Pajukanta

  294. Unidad de Biología Molecular y Medicina Genómica, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Mexico, Mexico

    Teresa Tusié-Luna

  295. Instituto de Investigaciones Biomédicas, UNAM, Ciudad de México, Mexico, Mexico

    Teresa Tusié-Luna

  296. Unidad de Biología Molecular y Medicina Genómica, Instituto de Investigaciones Bimédicas UNAM/ Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Mexico, Mexico

    Teresa Tusié-Luna

  297. Departamento de Endocrinología y Metabolismo, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Mexico, Mexico

    Carlos A. Aguilar-Salinas

  298. Dirección de Nutrición and Unidad de Estudios de Enfermedades Metabólicas, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Mexico, Mexico

    Carlos A. Aguilar-Salinas

  299. Department of Nutrition, Gillings School of Global Public Health, University of North Carolina, Chapel Hill, NC, USA

    Linda S. Adair & Penny Gordon-Larsen

  300. Carolina Population Center, University of North Carolina, Chapel Hill, NC, USA

    Linda S. Adair & Penny Gordon-Larsen

  301. USC–Office of Population Studies Foundation, University of San Carlos, Cebu City, Philippines

    Sonny Augustin Bechayda

  302. Department of Anthropology, Sociology, and History, University of San Carlos, Cebu City, Philippines

    Sonny Augustin Bechayda

  303. Department of Medicine, Faculty of Medicine, University of Kelaniya, Ragama, Sri Lanka

    H. Janaka de Silva

  304. Department of Public Health, Faculty of Medicine, University of Kelaniya, Ragama, Sri Lanka

    Ananda R. Wickremasinghe

  305. Departments of Pediatrics and Medicine, University of California, San Francisco, San Francisco, CA, USA

    Ronald M. Krauss

  306. Institute of Biomedical Sciences, Academia Sinica, Taipei, Taiwan

    Jer-Yuarn Wu

  307. Systems Genomics Laboratory, School of Biotechnology, Jawaharlal Nehru University, New Delhi, India

    Dwaipayan Bharadwaj

  308. Department of Medicine, University of Mississippi Medical Center, Jackson, MS, USA

    Adolfo Correa

  309. Department of Physiology and Biophysics, University of Mississippi Medical Center, Jackson, MS, USA

    James G. Wilson

  310. Department of Medical Sciences, Uppsala University, Uppsala, Sweden

    Lars Lind

  311. Department of Paediatrics, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore

    Chew-Kiat Heng

  312. Khoo Teck Puat–National University Children’s Medical Institute, National University Health System, Singapore, Singapore

    Chew-Kiat Heng

  313. Department of Medicine, University of North Carolina, Chapel Hill, NC, USA

    Amanda E. Nelson

  314. Department of Epidemiology, Gillings School of Global Public Health, University of North Carolina, Chapel Hill, NC, USA

    Yvonne M. Golightly

  315. Injury Prevention Research Center, University of North Carolina, Chapel Hill, NC, USA

    Yvonne M. Golightly

  316. Division of Physical Therapy, University of North Carolina, Chapel Hill, NC, USA

    Yvonne M. Golightly

  317. Department of Psychiatry, Amsterdam UMC, Vrije Universiteit Amsterdam, Amsterdam, The Netherlands

    Brenda Penninx

  318. Amsterdam Public Health Research Institute, VU Medical Center Amsterdam, Amsterdam, The Netherlands

    Brenda Penninx & Eco J. C. de Geus

  319. Department of Biochemistry, College of Medicine, Ewha Womans University, Seoul, Korea

    Hyung-Lae Kim

  320. Faculty of Health and Medicine, University of Newcastle, Newcastle, New South Wales, Australia

    John Attia & Rodney J. Scott

  321. Division of Biostatistics, Washington University School of Medicine, St Louis, MO, USA

    D. C. Rao

  322. University of Kentucky, College of Public Health, Lexington, KY, USA

    Donna K. Arnett

  323. Institute of Cellular Medicine (Diabetes), The Medical School, Newcastle University, Newcastle upon Tyne, UK

    Mark Walker

  324. Department of Population Health, Finnish Institute for Health and Welfare, Helsinki, Finland

    Heikki A. Koistinen

  325. University of Helsinki and Department of Medicine, Helsinki University Hospital, Helsinki, Finland

    Heikki A. Koistinen

  326. Minerva Foundation Institute for Medical Research, Helsinki, Finland

    Heikki A. Koistinen

  327. Academy of Scientific and Innovative Research (AcSIR), New Delhi, India

    Giriraj R. Chandak

  328. Diabetology Research Centre, KEM Hospital and Research Centre, Pune, India

    Chittaranjan S. Yajnik

  329. Programs in Metabolism and Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA

    Josep M. Mercader

  330. Diabetes Unit and Center for Genomic Medicine, Massachusetts General Hospital, Boston, MA, USA

    Josep M. Mercader

  331. Instituto Nacional de Salud Publica y Centro de Estudios en Diabetes, Cuernavaca, Mexico

    Clicerio Gonzalez Villalpando

  332. Instituto Nacional de Medicina Genómica, Mexico, Mexico

    Lorena Orozco

  333. Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, TX, USA

    Myriam Fornage

  334. Yong Loo Lin School of Medicine, National University of Singapore and National University Health System, Singapore, Singapore

    E. Shyong Tai & Rob M. van Dam

  335. Kurume University School of Medicine, Kurume, Japan

    Mitsuhiro Yokota

  336. Genetics, Merck Sharp & Dohme, Kenilworth, NJ, USA

    Dermot F. Reilly

  337. Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford, Oxford, UK

    Mark I. McCarthy

  338. Population Health and Genomics, University of Dundee, Ninwells Hospital and Medical School, Dundee, UK

    Colin N. A. Palmer

  339. Intramural Research Program, National Institute on Aging, Baltimore, MD, USA

    Eleanor Simonsick

  340. The Eye Hospital, School of Ophthalmology and Optometry, Wenzhou Medical University, Wenzhou, China

    Fan Lu & Jia Qu

  341. Shanghai Institute of Nutrition and Health, University of Chinese Academy of Sciences, Chinese Academy of Sciences, Shanghai, China

    Shaofeng Huo, Huaixing Li & Xu Lin

  342. Synlab Academy, Synlab, Mannheim, Germany

    Winfried März

  343. Clinical Institute of Medical and Chemical Laboratory Diagnostics, Medical University of Graz, Graz, Austria

    Winfried März

  344. Unidad de Investigacion Medica en Bioquimica, Hospital de Especialidades, Centro Medico Nacional Siglo XXI, Instituto Mexicano del Seguro Social, Mexico, Mexico

    Miguel Cruz

  345. Faculty of Medicine, University of Iceland, Reykjavik, Iceland

    Vilmundur Gudnason, Unnur Thorsteinsdottir & Kari Stefansson

  346. Department of Medicine, Faculty of Medecine, Université de Montréal, Quebec, Quebec, Canada

    Jean-Claude Tardif

  347. Department of Medicine, Faculty of Medicine, Université de Montréal, Montreal, Quebec, Canada

    Guillaume Lettre

  348. Department of Biomedical Data Sciences, Section Molecular Epidemiology, Leiden University Medical Center, Leiden, The Netherlands

    Leen M. ’t Hart

  349. Amsterdam UMC, Department of General Practice and Elderly Care, Amsterdam Public Health Research Institute, Amsterdam, The Netherlands

    Petra J. M. Elders

  350. Department of Genetics, University of Pennsylvania, Philadelphia, PA, USA

    Struan F. A. Grant & Daniel J. Rader

  351. Department of Medicine, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA

    Daniel J. Rader & Kyong-Mi Chang

  352. Department of Surgery, University of Pennsylvania, Philadelphia, PA, USA

    Scott M. Damrauer

  353. Corporal Michael Crescenz VA Medical Center, Philadelphia, PA, USA

    Scott M. Damrauer

  354. Institute of Social and Economic Research, University of Essex, Essex, UK

    Meena Kumari

  355. The Mindich Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, New York, NY, USA

    Ruth J. F. Loos

  356. Cardiovascular Health Research Unit, Departments of Medicine, Epidemiology and Health Services, University of Washington, Seattle, WA, USA

    Bruce M. Psaty

  357. Kaiser Permanent Washington Health Research Institute, Seattle, WA, USA

    Bruce M. Psaty

  358. Institute of Regional Health Research, University of Southern Denmark, Odense, Denmark

    Ivan Brandslund

  359. Danish Aging Research Center, University of Southern Denmark, Odense, Denmark

    Kaare Christensen

  360. Public Health, Faculty of Medicine, University of Helsinki, Helsinki, Finland

    Samuli Ripatti

  361. Broad Institute of MIT and Harvard, Cambridge, MA, USA

    Samuli Ripatti

  362. Center for Applied Genomics, Children’s Hospital of Philadelphia, Philadelphia, PA, USA

    Hakon Hakonarson

  363. Department of Pediatrics, Perelman School of Medicine, The University of Pennsylvania, Philadelphia, PA, USA

    Hakon Hakonarson & Struan F. A. Grant

  364. Division of Human Genetics, Children’s Hospital of Philadelphia, Philadelphia, PA, USA

    Struan F. A. Grant

  365. Institute of Genetic Epidemiology, Department of Genetics and Pharmacology, Medical University of Innsbruck, Innsbruck, Austria

    Florian Kronenberg

  366. School of Medicine, Southern University of Science and Technology, Shenzhen, China

    Dongfeng Gu

  367. Institute for Cardiogenetics, University of Lübeck, DZHK (German Research Centre for Cardiovascular Research), Partner site Hamburg/Lübeck/Kiel, and University Heart Center Lübeck, Lübeck, Germany

    Jeanette Erdmann

  368. Deutsches Herzzentrum München, Klinik für Herz- und Kreislauferkrankungen, Technische Universität München, Munich, Germany

    Heribert Schunkert

  369. Deutsches Zentrum für Herz-Kreislauf-Forschung (DZHK) e.V., Partner site Munich Heart Alliance, Munich, Germany

    Heribert Schunkert

  370. Department of Clinical Medicine, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

    Allan Linneberg

  371. Netherlands Heart Institute, Utrecht, The Netherlands

    J. Wouter Jukema

  372. Division of Cardiology, Department of Medicine, Massachusetts General Hospital, Boston, MA, USA

    Amit V. Khera

  373. Program of Medical and Population Genetics, Broad Institute, Cambridge, MA, USA

    Amit V. Khera & Sekar Kathiresan

  374. Center for Genomic Medicine, Massachusetts General Hospital, Boston, MA, USA

    Amit V. Khera

  375. Department of Medicine, Harvard Medical School, Boston, MA, USA

    Amit V. Khera & Sekar Kathiresan

  376. Northern Finland Birth Cohorts, Infrastructure for Population Studies, Faculty of Medicine, University of Oulu, Oulu, Finland

    Minna Männikkö

  377. Center for Life Course Health Research, Faculty of Medicine, University of Oulu, Oulu, Finland

    Marjo-Riitta Jarvelin

  378. Biocenter of Oulu, University of Oulu, Oulu, Finland

    Marjo-Riitta Jarvelin

  379. University Center for Primary Care and Public Health, Lausanne, Switzerland

    Zoltan Kutalik

  380. Institute for Genetic and Biomedical Research, Italian National Council of Research (IRGB CNR), Cagliari, Italy

    Francesco Cucca

  381. University of Sassari, Sassari, Italy

    Francesco Cucca

  382. Department of Clinical Epidemiology, Leiden University Medical Center, Leiden, The Netherlands

    Dennis O. Mook-Kanamori

  383. Department of Public Health and Primary Care, Leiden University Medical Center, Leiden, The Netherlands

    Dennis O. Mook-Kanamori

  384. Department of Internal Medicine, Division of Endocrinology, Leiden University Medical Center, Leiden, the Netherlands

    Ko Willems van Dijk

  385. Einthoven Laboratory for Experimental Vascular Medicine, Leiden University Medical Center, Leiden, the Netherlands

    Ko Willems van Dijk

  386. Department of Human Genetics, Leiden University Medical Center, Leiden, The Netherlands

    Ko Willems van Dijk

  387. Division of Cardiovascular Medicine, Radcliffe Department of Medicine, John Radcliffe Hospital, University of Oxford, Oxford, UK

    Hugh Watkins

  388. Population Health Research Institute, St George’s University of London, London, UK

    David P. Strachan

  389. National Heart and Lung Institute, Imperial College London, London, UK

    Peter Sever

  390. School of Public Health, Imperial College London, London, UK

    Neil Poulter

  391. OCDEM, University of Oxford, Churchill Hospital, Oxford, UK

    Fredrik Karpe & Matt J. Neville

  392. NIHR Oxford Biomedical Research Centre, Churchill Hospital, Oxford, UK

    Fredrik Karpe & Matt J. Neville

  393. Ophthalmology and Visual Sciences Academic Clinical Program (Eye ACP), Duke–NUS Medical School, Singapore, Singapore

    Ching-Yu Cheng, Tien-Yin Wong & Charumathi Sabanayagam

  394. DZHK (German Centre for Cardiovascular Research), Munich Heart Alliance Partner Site, Munich, Germany

    Annette Peters

  395. University of Exeter Medical School, University of Exeter, Exeter, UK

    Andrew T. Hattersley

  396. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden

    Nancy L. Pedersen & Patrik K. E. Magnusson

  397. Amsterdam Public Health, VU Medical Center Amsterdam, Amsterdam, The Netherlands

    Dorret I. Boomsma

  398. Amsterdam Reproduction and Development Research Institute, VU Medical Center Amsterdam, Amsterdam, the Netherlands

    Dorret I. Boomsma

  399. Framingham Heart Study, National Heart, Lung, and Blood Institute, US National Institutes of Health, Bethesda, MD, USA

    L. Adrienne Cupples

  400. Department of Genetics, School of Medicine, Mashhad University of Medical Sciences, Mashhad, Iran

    Mohsen Ghanbari

  401. Department of Genetics, Shanghai-MOST Key Laboratory of Health and Disease Genomics, Chinese National Human Genome Center at Shanghai, Shanghai, China

    Wei Huang

  402. TUM School of Medicine, Technical University of Munich (TUM) and Klinikum Rechts der Isar, Munich, Germany

    Eleftheria Zeggini

  403. Institute of Clinical Medicine, Internal Medicine, University of Eastern Finland and Kuopio University Hospital, Kuopio, Finland

    Johanna Kuusisto & Markku Laakso

  404. Stanford Cardiovascular Institute, Stanford University, Stanford, CA, USA

    Erik Ingelsson

  405. Stanford Diabetes Research Center, Stanford University, Stanford, CA, USA

    Erik Ingelsson

  406. Regeneron Pharmaceuticals, Tarrytown, NY, USA

    Goncalo Abecasis & Aris Baras

  407. Lee Kong Chian School of Medicine, Nanyang Technological University, Singapore, Singapore

    John C. Chambers

  408. Imperial College Healthcare NHS Trust, Imperial College London, London, UK

    John C. Chambers

  409. MRC–PHE Centre for Environment and Health, Imperial College London, London, UK

    Jaspal S. Kooner

  410. National Heart and Lung Institute, Imperial College London, London, UK

    Jaspal S. Kooner

  411. Institute for Minority Health Research, University of Illinois College of Medicine, Chicago, IL, USA

    Martha Daviglus

  412. Department of Biostatistics, Harvard T. H. Chan School of Public Health, Boston, MA, USA

    Peter Kraft

  413. QIMR Berghofer Medical Research Institute, Brisbane, Queensland, Australia

    Nicholas G. Martin & John B. Whitfield

  414. Faisalabad Institute of Cardiology, Faislabad, Pakistan

    Shahid Abbas

  415. Department of Medicine, Columbia University Irving Medical Center, New York, NY, USA

    Danish Saleheen

  416. Department of Cardiology, Columbia University Irving Medical Center, New York, NY, USA

    Danish Saleheen

  417. Big Data Institute, University of Oxford, Oxford, UK

    Robin G. Walters

  418. National Institute for Health Research Oxford Biomedical Research Centre, Oxford University Hospitals, Oxford, UK

    Michael V. Holmes

  419. Aberdeen Centre for Health Data Science, School of Medicine, Medical Science and Nutrition, University of Aberdeen, Aberdeen, UK

    Corri Black

  420. Division of Population Health and Genomics, Ninewells Hospital and Medical School, University of Dundee, Dundee, UK

    Blair H. Smith

  421. Biomedical and Translational Informatics, Geisinger Health, Danville, PA, USA

    Anne E. Justice

  422. Department of Biomedical Informatics, Vanderbilt University Medical Center, Nashville, TN, USA

    Wei-Qi Wei

  423. Departments of Medicine (Medical Genetics) and Genome Sciences, University of Washington Medical Center, Seattle, WA, USA

    Gail P. Jarvik

  424. Center for Autoimmune Genomics and Etiology, Cincinnati Children’s Hospital Medical Center (CCHMC), Cincinnati, OH, USA

    Bahram Namjou

  425. Division of Endocrinology, Metabolism, and Molecular Medicine, Department of Medicine, Northwestern University, Feinberg School of Medicine, Chicago, IL, USA

    M. Geoffrey Hayes

  426. Department of Anthropology, Northwestern University, Evanston, IL, USA

    M. Geoffrey Hayes

  427. Center for Genetic Medicine, Northwestern University, Feinberg School of Medicine, Chicago, IL, USA

    M. Geoffrey Hayes

  428. HUNT Research Centre, Department of Public Health and Nursing, NTNU, Norwegian University of Science and Technology, Levanger, Norway

    Kristian Hveem & Bjørn Olav Åsvold

  429. Department of Medicine, Levanger Hospital, Nord-Trøndelag Hospital Trust, Levanger, Norway

    Kristian Hveem

  430. Department of Endocrinology, St Olavs Hospital, Trondheim University Hospital, Trondheim, Norway

    Bjørn Olav Åsvold

  431. RIKEN Center for Integrative Medical Sciences, Yokohama, Japan

    Michiaki Kubo

  432. Laboratory of Complex Trait Genomics, Department of Computational Biology and Medical Sciences, Graduate School of Frontier Sciences, The University of Tokyo, Tokyo, Japan

    Yoichiro Kamatani

  433. Laboratory of Statistical Immunology, WPI Immunology Frontier Research Center, Osaka University, Osaka, Japan

    Yukinori Okada

  434. Integrated Frontier Research for Medical Science Division, Institute for Open and Transdisciplinary Research Initiatives, Osaka University, Osaka, Japan

    Yukinori Okada

  435. Division of Molecular Pathology, Institute of Medical Science, The University of Tokyo, Tokyo, Japan

    Yoshinori Murakami

  436. VA Boston Healthcare System, Boston, MA, USA

    Yuk-Lam Ho, Kelly Cho, Christopher J. O’Donnell & John M. Gaziano

  437. VA Informatics and Computing Infrastructure, VA Salt Lake City Health Care System, Salt Lake City, UT, USA

    Julie A. Lynch

  438. University of Massachusetts, Boston, MA, USA

    Julie A. Lynch

  439. Cardiovascular Institute, Stanford University School of Medicine, Stanford, CA, USA

    Philip S. Tsao & Themistocles L. Assimes

  440. Corporal Michael J. Crescenz VA Medical Center, Philadelphia, PA, USA

    Kyong-Mi Chang

  441. Department of Medicine, Brigham Women’s Hospital, Boston, MA, USA

    Kelly Cho, Christopher J. O’Donnell & John M. Gaziano

  442. Atlanta VA Medical Center, Atlanta, GA, USA

    Peter Wilson

  443. Division of Cardiology, Emory University School of Medicine, Atlanta, GA, USA

    Peter Wilson

  444. School of Electrical and Information Engineering, University of the Witwatersrand, Johannesburg, South Africa

    Scott Hazelhurst

  445. Division of Human Genetics, National Health Laboratory Service and School of Pathology, Faculty of Health Sciences, University of the Witwatersrand, Johannesburg, South Africa

    Michèle Ramsay

  446. School of Basic and Medical Biosciences, Faculty of Life Sciences and Medicine, King’s College London, London, UK

    Richard C. Trembath

  447. Departments of Pediatrics and Genetics, Harvard Medical School, Boston, MA, USA

    Joel N. Hirschhorn

  448. Center for Genomic Medicine, Department of Medicine, Massachusetts General Hospital, Boston, MA, USA

    Sekar Kathiresan

  449. Cardiology Division, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA

    Pradeep Natarajan

  450. Department of Medicine, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA

    Pradeep Natarajan

  451. Cardiovascular Research Center and Center for Genomic Medicine, Massachusetts General Hospital, Boston, MA, USA

    Pradeep Natarajan

  452. Centre for Genetics and Genomics Versus Arthritis, Centre for Musculoskeletal Research, Division of Musculoskeletal and Dermatological Sciences, The University of Manchester, Manchester, UK

    Andrew P. Morris

  453. Princess Al-Jawhara Al-Brahim Centre of Excellence in Research of Hereditary Disorders (PACER-HD), King Abdulaziz University, Jeddah, Saudi Arabia

    Panos Deloukas & Cristen J. Willer

  454. Department of Human Genetics, University of Michigan, Ann Arbor, MI, USA

    Cristen J. Willer

Authors
  1. Sarah E. Graham
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  2. Shoa L. Clarke
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  3. Kuan-Han H. Wu
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